RSPB 1933 0071

64
612.492.5
The Effect of Hypophysectomy of the Rabbit.
By W illiam E. W hite , Cutting Travelling Fellow, Columbia University.

(From the National Institute for Medical Research, London.)
Downloaded from https://royalsocietypublishing.org/ on 26 October 2022
(Communicated by A. S. Parkes, F.R.S.—Received July 13, 1933.)
[Plates 2-5.]
I. Introduction.
In recent years the effect of surgically removing the pituitary gland has
received considerable attention. Two classes of vertebrates, Amphibia and
Mammalia, have largely been used in these studies. Practically all the
experimental work on ablation in mammals has been done on two species,
namely, dogs (Cushing et ol., 1910 ; Aschner, 1912 ; Houssay, 1931 ; and many
others), and more recently rats (Smith, 1930 ; and others).* Following the
epoch-making discovery of Smith and Engle (1926) and Aschheim and Zondek
(1927), that implanted pituitary gland will hasten sexual maturity in the rat,
a number of workers have been attempting to isolate the hormones from the
anterior lobe. It has become increasingly apparent that adequate test animals
are necessary!for the physiological assay of the various hormone fractions and
it is now equally apparent that only with animals whose pituitaries have been
removed may purified products be assayed with any certainty.
The peculiarities of the rabbit’s reproductive cycle and also its possible
utility for hormone assay, make it unusually interesting for studies on the
effect of hypophysectomy, but the inaccessibility of the gland has somewhat
retarded work on this species. In conjunction with Dr. P. E. Smith, however,
method for ablating the pituitary was recently elaborated (Smith and White,
1931) which left the animals essentially healthy so that they survived for
considerable periods. I t therefore seemed interesting to determine the extent
of the changes caused by hypophysectomy in the rabbit and to attempt
substitution therapy with purified hormones. The data on untreated hypo-
physectomized animals will be presented in the present paper.
* For an adequate review of the innumerable papers which have appeared on this
subject see Cushing (1912) and Smith (1930) among others.
Effect of Hypophysectomy of the Rabbit. 65
II. Methods.
Since the rabbits used in these experiments were purchased from dealers it
was not possible to obtain litter-mate controls. Nevertheless, the animals
were all normal mature individuals, well fed and seemingly in good condition.
They were kept in quarantine for approximately one month before use and
were then removed to a room maintained a t a constant temperature. It is
highly desirable for hypophysectomized animals to be protected from sudden
changes in temperature as their resistance is considerably below that of normal

individuals. A temperature around 75° F. seems to be optimum.
Operative Technique.—The topography of the rabbit skull makes access to
the pituitary fairly difficult. I t is essential when complete hypophysectomy
is attempted to have the gland clearly visible. Partial removal sometimes
occurs even with an unobstructed view of the gland, and it is indeed optimistic
to attempt complete removal by a blind operation.
The size and shape of the mandible makes a parapharyngeal approach
impracticable in these species. Accordingly, a transbuccal approach, first
successfully used by Aschner in the dog, was attempted. Some difficulty was
experienced in getting the animal properly orientated for accurate operating.
Finally, a special head holder was devised. This apparatus has proved so
helpful that it will be briefly described. The animal is anaesthetized, the mouth
opened, and the jaws held sufficiently rigid by cords attached to the upper and
lower incisors. The tongue is depressed by a suitable retractor which allows
a clear view of the operative field without causing any cyanosis of the tongue.
The head is held perpendicular by two adjustable side-plates. The device is
constructed to allow of additional orientation without removing or soiling the
sterile cloth which covers the board at this stage. Since mucus tends to
collect in the nasal passages when the animal is left under urethane or other
protracted anaesthesia, it is highly desirable for the animal to regain con
sciousness as soon as the operation is completed. Ether anaesthesia proved
satisfactory. A mixture of oxygen and 5% C02, when properly regulated
and bubbled through ether, and then conveyed into the mouth cavity by a
tube at the side of the tongue retractor, has given very even and easily regulated
anaesthesia. No tracheotomy is necessary and the animal’s normal breathing
can readily be watched by an occasional glance a t the chest, and checked by
attention to the colour of the blood vessels and so forth. This apparatus has
made possible a single-handed operation and has been used by the writer
without assistance for about 18 months.
VOL. cxiv.—B. F
66 W. E. White.
When the animal is properly orientated, a median incision, about l | cm.

in length, is made in the soft palate, fig. 1. The injection of a few drops of
adrenalin and cocaine solution prevents any slight haemorrhage. The incision
is then opened and the margins held securely by a small spring retractor which
is inserted with the aid of a pair of grooved Spencer-Wells forceps. The nasal
mucosa is then compressed until it is devascularized. When this is accom-
Retractor
Tube for ether
Tongue
Site of incision
Soft palate
F ig. 1.—General view of the mouth of the rabbit, showing topography of the soft palate
and the site and extent of the incision in the oral mucosa.
plished the cavernous foramen in the basi-sphenoid becomes visible and is the
landmark for the later stages, fig. 2.
In earlier work a small quantity (0-1 c.c.) of wax warmed to just above the
melting point was injected into this foramen. In all the later work, however,
cold wax has been pressed into the foramen, since heating the wax necessitated
handling of syringes, the use of alcohol lamps, and other chances of soiling the
instruments. When the cavernous sinus is occluded with wax, an elliptical
hole is drilled in the sphenoid bone, cephalic to the cavernous foramen which
forms the posterior margin of the hole. An ordinary dental reamer with an
electric foot switch is very useful for this purpose. The hole should be about
5-6 mm. long and about 2 mm. across, or as wide as can be made with safety,
since haemorrhage becomes troublesome if the lateral sinuses are opened,
fig. 3. The wax is then removed from the ventral surface of the pituitary
which is covered with marrow and often with a very fragile layer of bone. The
pituitary capsule is then slit with a sharp iridectomy knife and the entire
contents removed with appropriate cannulse and negative pressure. This is
Cavernous foramen as
- seen through mucosa
Compressed nasal
mucosa
—Margin of oral
I? v - J I mucosa
----- Spring retractor
F ig. 2.— Exposed nasal cavity, showing cavernous foramen as seen through the com
pressed nasal mucosa.
not a blind operation; all essential structures are clearly visible. There is
little or no haemorrhage, and any slight seepage readily clots. The hole is
then sealed with bone wax, the nasal mucosa swabbed with antiseptic solution,
and the incision in the oral mucosa sutured with silk, which sloughs out as the
wound heals.
The operation is admittedly through a septic field. The procedure, how
ever, is carried out under as favourable conditions as possible. The operating
cloths and instruments are sterilized and the mouth cavity swabbed with a
68 W. E. White.
5% solution of acriflavine before the first incision is made. Likewise

the nasal cavity is cleaned when entered and all reasonable precautions taken.
There is little trouble from infected wounds. Animals have been killed at
intervals up to 9 weeks after operation, at which time they were in excellent
condition and would doubtless have survived for much longer periods.
The technique is somewhat lengthy. Several operations have been per
formed in 45 minutes, but about an hour is the usual time. By far the most
unsatisfactory stage is the compression of the nasal mucosa which takes about
^Sphenoid bone and posterior

s' margin of cavernous fora-
\ men
_ - ' Pituitary gland
---- Nasal mucosa
- Edge of oral mucosa
F ig. 3.—Later stage, showing nasal mucosa slit, basi-sphenoid drilled and pituitary
capsule exposed.
half the total time required for pituitary ablations by this method. The
procedure, however, is necessary in this essentially bloodless operation. A
shorter and very ingenious method for pituitary ablation has recently been
described by Firor (1933) who employs an intra-orbital approach. The
operation requires much less time, but the gland is, apparently, only im
perfectly visible by this approach.
The completeness of the pituitary removal was always checked by serial
sections of the contents of the sella turcica, and in the more critical cases the
entire pituitary region was sectioned. In a few cases of obvious partial
removal, the residual gland was removed and weighed, and later sectioned to
confirm the diagnosis. A number of animals had only microscopical fragments
of gland and physiologically resembled completely hypophysectomized indi
viduals, but the present report includes only animals in which pituitary tissue
was completely absent.
Histological Technique.—All organs were weighed fresh, and were then fixed
in Bouin’s fluid (except for the adrenals, see p. 76), cut at 7 [x or 5 jx, and
stained in haemalum and eosin.
Control Material.—Control tissues were obtained from 10 normal female

rabbits and 5 normal males. In each plate, illustrations of the normal organs
are shown alongside those of the organs after hypophysectomy.
Treatment of Results.—The statistical examination of the weights of the
different organs is complicated by the fact that the animals were killed at
varying times after hypophysectomy. Except for the testes, however, there
is no evidence of regular progressive decrease in size, and it has therefore
seemed legitimate to take the average for the group. In calculating the weight
of an organ as per cent, body weight, the gross weight of the animal a t
autopsy has been used. In the females, the average body weight of the
operated animals is so near that of the controls, that calculations of weight of
an organ as per cent, body weight have not been included.
III. General Effects of Hypophysectomy.

Post-operative Complications.—Adult rabbits survive total ablation of the
pituitary gland fairly well. The most frequent complication after the opera
tion has been a respiratory infection, popularly known as csnuffles,’ in which a
purulent exudate is discharged from the nose. The animal rooms used
during the past year, however, have been quite free from the causative organism,
and infections of any sort have been rare. Occasionally, when the cannula is
inadvertently applied to the pituitary stalk while aspirating the gland, varying
amounts of cerebro-spinal fluid escape. If any considerable quantity is lost
death generally follows within 24 hours, although the animal may eat and
appear to recover from the immediate effects of the operation.
Spontaneous Hypoglyccemia.—A few animals which had survived for 2 or
3 weeks after hypophysectomy, and which were apparently healthy 10-12
hours earlier, were found in a moribund condition. Blood sugar determinations
showed these animals to be in a severe hypoglycaemic coma. This spontaneous
flyp°glyc8emia even when food had not been withdrawn was surprising, and
led to a study of carbohydrate metabolism in the hypophysectomized rabbit.
70 W. E. White.
Various aspects of this work have been undertaken in collaboration with Dr.
A. B. Corkill and Mr. H. P. Marks, and will be discussed in another paper.
I t may be mentioned, however, that 12 hours was the maximum fast which
could be allowed with safety, as several animals died after 24 hours without food.
Behaviour of Animals after Operation.—Twenty-four hours after operation
hypophysectomized rabbits are essentially recovered and closely resemble
normal animals in general appearance and behaviour. As the time after
operation increases they become slightly sluggish. The body temperature
averages around 100° F., whereas normal control animals have body tempera
tures ranging around 102° F., when similarly tested. There is a striking loss
of sexual interest after operation. This is particularly well shown in HR 17
which had been used for breeding before operation and was considered to be
the best breeding male in the laboratory. After hypophysectomy it tried
to copulate only once, although put with several cestrous females after varying
intervals. This single attempt was completely unsuccessful, although the
female readily copulated when placed with a normal male. No other males, nor
any females, have shown any interest in the opposite sex.
Fat Deposition.—I t has frequently been reported that animals tend to deposit
fat after pituitary ablation. This now seems to be related to injury of the
hypothalamus rather than to a pituitary deficiency (see Smith, 1930). In
the rabbit, brain injury is improbable from a ventral approach, as there is a
layer of bone completely separating the pituitary from the brain, and this
species has not shown any appreciable deposition of fat for at least 9 weeks
after hypophysectomy, the longest stage investigated. There is on the
other hand always a definite, though variable decrease in body weight after
operation.
Loss of Hair.—After hypophysectomy there seems to be at least a partial
shedding of hair. In most varieties of rabbits this shedding is not pronounced.
However, in three individuals of a blue Beveran strain, the coat became quite
shaggy about a fortnight after operation, although the animals otherwise
seemed to be in good condition. It is not known whether this is a specific
effect of hypophysectomy or not, but certainly it is a general occurrence.
The most clearly marked changes in adult rabbits after hypophysectomy are
in the reproductive system, adrenal cortex, thyroids, and liver weight.
IV. The Male Reproductive System.

As has already been mentioned, there is an immediate and complete loss of
potency in the male rabbit after hypophysectomy. This physiological mani-
Effect of Hypophysectomy of the Rabbit.
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Body weight.
Days after
Animal. hypophy Thyroid Adrenals Ovaries Uterus Kidneys Liver Spleen
sectomy. At operation, At autopsy, weight, gm. weight, gm. weight, gm. weight, gm. weight, gm. weight, gm. weight, gm.
kg. kg.
HR 18 ....... 3*20 160

HE 22 ....... 42-0 0-44
o o
3*25 13-8 48-0 0*70
HE 2 .......... 410 18-5
HR 12 .......
HR 13 .......
t^CO^h h
2*60 120
HE 25 .......
W. E. White.
2-50 OOCO<N<NCOO 12- 5 40 0 0-85

HE 20 ....... 4*20 CO 18-5 60*0
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13- 5 62-0 0-60
Average weight............ 3-28 3-12 0-145 0-227 0-300 3-15
Average weight as per 15 0 50-4 0-65
cent. B.W............ 0-0047 0-007 0-009 0-101 0-481 1-6 0-02
festation is accompanied by a striking regression in both the testis and acces

sory organs.
Changes in the Testis.—The testes undergo a profound atrophy after ablation
of the anterior pituitary. Usually they decrease to about one-fifth normal
size, fig. 4, and become quite flabby. As a rule they do not ascend into the
abdominal cavity, but remain in the scrotum. Early changes appear 6 days
after operation, a few abnormal cells being found in the lumina of the semini
ferous tubules. After 11 days the tubules decrease somewhat in diameter
025
0 *20 '
&
CTDo1
~iO 20 50 40 50 60 70
Days after hypophysectomy
F ig. 4.—Scatter diagram of weight of testes at various times after hypophysectomy.
and regression is more evident. By 3 weeks after operation the testis is con
siderably altered in appearance. No spermatozoa are present; the cells in
the tubules are limited to spermatogonia and primary spermatocytes, a few
of which are still dividing. Mitotic figures are no longer found at 4 weeks,
figs. 5 and 6, Plate 2, the tubules now containing only spermatogonia, an
occasional spermatocyte, and a Sertoli reticulum. This condition is unchanged
o
for at least 67 days, the latest stage examined.
015
The proportion
iO of interstitial tissue increases in the testis after hypophy
sectomy. From rough quantitative estimations it is clear that the change
OJO
is only relative, due to the remarkable decrease in the size of the seminiferous
tubules, and is not a real hyperplasia.
Changes in the Accessory Organs.—The accessory organs undergo atrophy
comparable to that of the testis. The epididymis decreases both in size and
weight. Histologically, its tubular epithelium is much flattened and inactive.
&perm gradually disappear and are entirely absent after 9 weeks, fig. 8,
74 W. E. White.
Table III.—Male Rabbits. Effect of Hypophysectomy on the Weights of

various Organs.
Change after
Normal controls. Hypophysectomized. hypophysectomy
%
Average Average
Average as % Average as % As %
weight, weight As net
body body weight. body
gm. gm. weight.
weight. weight.
Body weight ............... 2600

2230 - 21*
Thyroid.......................... 0-221 0-0085 0-154 0-0069 - 30 - 19
Adrenals ...................... 0-308 0-0119 0-296 0-0133 - 4 + 12
Testes .......................... 5-1 0-196 — — —
Pituitary ...................... 0-033 0-0014 — — — —
Kidneys ...................... 16-1 0-62 13-6 0-61 - 15 - 1
Liver .......................... 76-2 2-93 41-0 1-84 - 45 - 37
Spleen .......................... 1-3 0-05 10 0-045 - 23 - 10
No. of animals ........... 5 9 — —
* See Table I.
Table IV.—Female Rabbits. Effect of Hypophysectomy on the Weights of

various Organs.
Change after
Normal controls. hypophy
HyP0Physectomized. sectomy
%
Average Average as Average Average as As net

weight, gm. body %. %. weight.
weight. weight, gm. body weight.
p
Body weight .............. 3200 _ 3120 _ - 5*
Thyroid.......................... 0-181 0-0056 0-145 0-0047 -2 0
Adrenals ...................... 0-603 0-018 0-227 0-007 -6 2
Ovaries ...................... 1-098 0-034 0-300 0-009 -7 2
Uterus .......................... 11-6 0-363 3-15 0-101 -7 0
Pituitary ...................... 0-064 0-002 — — —
Kidneys ...................... 20-2 0-632 15-00 0-481 -2 6
Liver .......................... 121-0 3-780 50-40 1-60 -5 8
Spleen .......................... 2-85 0-089 0-65 0-02 -7 7
No. of animals ........... 10 iS —
* See Table II.

Plate 2. Similar changes occur in the ductus deferens, fig. 15, Plate 4. The
lining epithelium, which is fairly high and often convoluted in the normal
organ, becomes flattened as in the epididymis. The lumen of the ductus is
filled with varying amounts of necrotic material for some time after operation.
The changes in the prostate and seminal vesicles are typically those which
occur following sudden atrophy of any secreting gland. Besides a decrease
in total weight there is a regression of the epithelial lining, affecting both the
cytoplasm and, to some extent, the nuclei of the cells.
V. The Female Reproductive System.

Effects on the Ovary.—The changes in the ovaries are striking and rapid in
appearance. Within 72 hours the larger follicles have decreased in size and
after 6 days only small follicles can be seen grossly. For several weeks follicles
with small antra can be found microscopically, but after this time only egg
nests and very small follicles undergoing atresia are seen, fig. 10, Plate 3.
The main body of the ovary, largely interstitial tissue in adult rabbits, appears
little changed histologically, but the ovary weight decreases to about one-third
its average for adults. Corpora lutea of pregnancy do not seem to persist for
such long periods as Smith (1930) found in the rat. In several animals, which
are not considered in the present data, but which were operated on at various
times during pregnancy, the corpora lutea were difficult to identify 3 to 5 weeks
after operation.
Changes in the TJterus and Fallopian Tube.—The uterus also undergoes a
very striking regression, figs. 11, 12, 14, Plate 3. There is a diminution not
only in the mucosa, but also in the muscular layers, and the whole uterus
resembles that of a castrate. By 3 weeks after operation the parous uterus
may weigh only 4-5 gm. and the non-parous considerably less. Occasionally
the lumen is filled with a clear fluid, but more often it is collapsed and in all
animals the uterus is very pale and flaccid. The diameter of the Fallopian
tube decreases until it is often difficult to find in the surrounding fat. The
epithelium changes from fairly high pseudo-stratified to a compact cuboidal
type and becomes less convoluted, figs. 17, 18, Plate 4.
Changes in Vagina.—The vagina undergoes changes after hypophysectomy
comparable to the regression in the uterus. In the normal oestrous female the
vagina is remarkably large. In sections the wall is generally convoluted and
lined by actively secreting mucous cells. In the operated animal the epithelium
loses all secretory activity and becomes little more than a layer of nuclei lining
the lumen, fig. 21, Plate 4.
76 W. E. White.
Effect on Pregnancy—In a previous paper (White, 1932) it was shown that

either hypophysectomy or blocking the pituitary circulation causes a termina
tion of pregnancy in the rabbit. Operations performed during the last half
of pregnancy invariably cause abortion within 48-72 hours. This work has
recently been confirmed by Firor (1933).
VI. The Adrenal Glands.

The adrenal weights in hypophysectomized rabbits are below the average

for normal adults, in the female markedly so, suggesting that the adrenals
regress after pituitary ablation in this species. Smith clearly showed that this
occurs in the rat. Histologically the main changes occur in the cortex, fig. 22,
Plate 5. The cells in all layers appear smaller in size so that their nuclei
are closer together. There is possibly also a slight decrease in the diameter
of the nuclei, figs. 24, 25, Plate 5. In the inner layer of the zona fasciculata
they tend to lose their strand-like arrangement.
Sometimes, though not always, haemorrhage is found in the juxta-medullary
zone. The proportion of cortex to medulla is somewhat difficult to determine
in the rabbit adrenal owing to the varied and irregular shape of the gland.
In most experiments half of a gland was fixed in Ciaccio’s solution, and the
other part in Zenker’s or Bouin’s fixative for staining in the ordinary way.
The Ciaccio fixed pieces were mounted on several slides, some of which were
examined unstained in glycerine and others stained with iron hematoxylin.
Histological examination of the sections showed that the medulla was un
changed and still showed the typical chromophile reaction. Physiological
and colorimetric analyses, which were kindly done by Dr. H. Schild, showed
that the amount of adrenaline per gram adrenal was approximately normal;
certainly not below, but possibly a little above, the average. This also
demonstrates that the medulla remains normal and that the loss in weight is
practically all due to cortical atrophy.
VII. The Thyroid Glands.

The thyroids apparently undergo only a very slight regression in adult
rabbits after hypophysectomy, as judged by total weight. The weight as
per cent, body weight for operated animals shows a decrease of about one-fifth.
It is difficult to dissect the fat from the thyroids, but all weighings were made
by the writer so that any error is probably constant for the two different series.
The histological changes are more clearly marked. The normal follicular
epithelium is of a fairly high cuboidal type, fig. 27, Plate 5. After hypophy
sectomy this epithelium becomes considerably flatter, sometimes approaching
a squamous type, fig. 26, Plate 5. This regression of the epithelium is un
doubtedly accompanied by a lowered thyroid activity, although the colloid
has the normal staining reaction and is found in about the same amounts as
in normal individuals. I t was not possible to obtain data on basal metabolism,
but the lower body temperature of operated animals suggests th at the basal
rate is probably low, owing partly to the decreased activity of the thyroid.
VIII. Changesin other Organs.
In hypophysectomized rabbits there is a definite loss in liver weight, though

this is not accompanied by any striking histological changes. Data on glycogen
content, as well as on insulin sensitivity and glucose tolerance, are considered in
another paper (Corkill, Marks and White) and will not be discussed here.
There may be a slight change in the parathyroids after hypophysectomy.
Any change is limited to a slight decrease in cell size. The external parathyroids
were not examined, so that the observations are restricted to the internal para
thyroids which are imbedded in the thyroid and were studied when this organ
was sectioned.
Weighings were also made of the kidneys and spleen. There is apparently
no significant change in the weight of either organ in the male. Both organs
in the female, especially the spleen, appear to decrease. Such results are to
be expected from data on a heterogeneous group where the loss in body weight
was often slight.
The writer wishes to express his gratitude to his former professor, Dr. P. E.
Smith, with whom the technique here described was worked out. He would
also like to express his appreciation to the Medical Research Council and to
Sir Henry Dale, Sec. R.S., for the hospitality of the National Institute for
Medical Research during the past year, and finally, he takes pleasure in
acknowledging the generous assistance of Dr. A. S. Parkes, F.R.S., in the
preparation of the manuscript and helpful criticisms during the course of
work in his laboratory.
IX. Summary.
Hypophysectomy in the rabbit is followed by :—
(1) A loss in body weight and a slight fall in body temperature.
78 Effect of Hypophysectomy of the Rabbit.
(2) A pronounced regression of both the male and female reproductive organs.
(3) A decrease in adrenal weight caused by atrophy of the cortex. This
regression seems to be merely a decrease in size of the cells and not in their
number.
(4) Changes in the structure of the thyroid accompanied by slight loss in
weight. The colloid is normal in amount, but the follicular epithelium much
flattened.
(5) A loss in the weight of the liver.
DESCRIPTION OF PLATES.
P late 2.
F ig . 5.—Testes of HR 19, 30 days after hypophysectomy. x 11.
F ig . 6.—Testes of HR 6, 42 days after hypophysectomy, showing histological details.
X 200. -
F ig. 7.—Normal rabbit testis for comparison with fig. 6. X 200.
F ig . 8.—Head of epididymis of HR 17, 67 days after hypophysectomy. x 35.
F ig . 9.—Normal epididymis, for comparison with fig. 8. x 35.
P late 3.
F ig . 10.—Ovary of HR 2, 30 days after hypophysectomy, showing absence of follicles.
X 8.
F ig . 11.—Uterus of HR 2, 30 days after hypophysectomy, parous animal. X 12.
F ig . 12.—Uterus of rabbit ovariectomized 6 months previously. X 12.
F ig . 13.—Normal cestrous uterus, for comparison with figs. 11 and 12. X 12.
F ig . 14.—Uterus of HR 13, 30 days after hypophysectomy, non-parous uterus. X 14.
P late 4.
F ig . 15.—Ductus deferens of HR 21, 26 days after hypophysectomy. X 35.
F ig . 16.—Normal ductus deferens for comparison with fig. 15. X 35.
F ig. 17.—Fallopian tube of HR 2, 30 days after hypophysectomy. See Plate 3 fig. 11.
X 28.
F ig. 18.—Fallopian tube of HR 13, 30 days after hypophysectomy, showing extreme
atrophy. X 28.
F ig . 19.—Normal Fallopian tube from cestrous rabbit for comparison with figs. 17 and 18.
X 28.
F ig . 20.—Normal vagina from cestrous female, showing active epithelium. X 35.
F ig . 21.—Vagina of HR 14, 67 days after hypophysectomy, showing atrophied epithelium.
X 35.
P late 5.
F ig . 22.—Adrenal of HR 5, 45 days after hypophysectomy (Ciaccio fixative), showing
normal chromophile reaction. X 14.
F ig . 23.—Normal adrenal for comparison with fig. 22. X 14.
F ig . 24.—Adrenal of HR 5, showing histological detail of practically whole width of cortex.
Ciaccio fixative, hsematoxylin stain. X 190.
White.
Proc. Roy.
(Facing p. 78 .)
Soc.,B, vol. 114, PI.
Downloaded from https://royalsocietypublishing.org/ on 26 October 2022 White.
Proc. Roy. Soc., B, vol. 114,
3.
Downloaded from https://royalsocietypublishing.org/ on 26 October 2022 White
Proc.
Roy. Soc.,B,
114. 4.
White. Proc. Roy. Soc., B, 114, PI.
-V 'X
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-
n • •
* * •*J* ******
7*
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; *? • <V..,~;.C24
o%
h Jk 1 w v
m * 'V ryY, ;/i
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Visual Purple 79
in the Mammalian Retina.
F ig . 25.—Normal adrenal for comparison with fig. 24, showing section taken midway
between medulla and capsule. Similar technique. X 190.
p IG 26.—Thyroid of HR 6, 42 days after hypophysectomy, showing flattened follicular
epithelium, x 190.
j?1Q' 27.—Normal thyroid, showing cuboidal epithelium. Compare fig. 26. X 190.
REFERENCES.
Ascheim and Zondek (1927). ‘ Arch. Gynak.’ vol. 130, p. 1.
Aschner (1912). 4Pfliiger’s Arch.,’ vol. 146, p. 7.
Crowes, Cushing and Hormans (1910). 4Bull. Johns Hopkins Hosp.,’ vol. 21, p. 127.
Cushing (1912). 44 The Pituitary Body and its Disorders.” Philadelphia, Lippincott.
Firor (1933). 4Amer. J. Physiol.,’ vol. 104, p. 204.
Houssay and Biasotti (1931). 4Pfliiger’s Arch.,’ vol. 227, p. 644 ; vol. 239, p. 651.
Smith (1930). 4Amer. J. Anat.,’ vol. 45, p. 205.
Smith and Engle (1926). ‘ Proc. Soc. Exp. Biol. N. Y .,’ vol. 24, p. 131.
Smith and White (1931). 4J. Amer. Med. Assoc.,’ vol. 97, p. 1861.
White (1932). 4Amer. J. Physiol.,’ vol. 102, p. 505.
6 l 2 .843.14
Factors affecting the Development and Regeneration of Visual Purple
in the Mammalian Retina.
By K atharine Tansley ,* The Strangeways Research Laboratory, Cambridge,
and The Department of Physiology, University College, London.
(Communicated by C. E.Lovatt Evans, F.R.S.—Received August 21, 1933.)
[P lates 6 a n d 7.]
Introduction.
The duplicity theory of vision ascribes the sensitivity of the eye to illumina
tions below 0*01 f.c. to the bleaching of the visual purple contained in the rods
of the retina and the phenomenon of dark adaptation is explained as being
due to the regeneration of this substance. It has been shown that there is a
close correlation between the rate of regeneration of visual purple and the rate
of dark adaptation and that conditions which affect one will also affect the
other (Tansley, 1931).
Hecht (1920) by mathematical reasoning from the curves of dark adaptation
in man has come to the conclusion that the regeneration of visual purple
* Working with a part-time grant from the Medical Research Council.

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64

By W illiam E. W hite , Cutting Travelling Fellow, Columbia University.

(Communicated by A. S. Parkes, F.R.S.—Received July 13, 1933.)

changes in temperature as their resistance is considerably below that of normal

When the animal is properly orientated, a median incision, about l | cm.

----- Spring retractor

5% solution of acriflavine before the first incision is made. Likewise

^Sphenoid bone and posterior

_ - ' Pituitary gland

---- Nasal mucosa

- Edge of oral mucosa

Control Material.—Control tissues were obtained from 10 normal female

III. General Effects of Hypophysectomy.

IV. The Male Reproductive System.

HR 18 ....... 3*20 160

2-50 OOCO<N<NCOO 12- 5 40 0 0-85

festation is accompanied by a striking regression in both the testis and acces­

Table III.—Male Rabbits. Effect of Hypophysectomy on the Weights of

Body weight ............... 2600

Table IV.—Female Rabbits. Effect of Hypophysectomy on the Weights of

Average Average as Average Average as As net

No. of animals ........... 10 iS —

* See Table II.

V. The Female Reproductive System.

Effect on Pregnancy—In a previous paper (White, 1932) it was shown that

VI. The Adrenal Glands.

The adrenal weights in hypophysectomized rabbits are below the average

VII. The Thyroid Glands.

VIII. Changesin other Organs.

In hypophysectomized rabbits there is a definite loss in liver weight, though

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festation is accompanied by a striking regression in both the testis and acces